Chiang Mai Journal of Science

Updated Taxonomic Insights into Auricularia (Auriculariales, Basidiomycota) in Thailand, with a New Record of Auricularia sinodelicata

1. INTRODUCTION

     The genus Auricularia was first proposed in 1780 with A. mesenterica as the type species [1]. This genus belongs to the family Auriculariaceae, order Auriculariales, and the phylum Basidiomycota [2–4]. Morphologically, Auricularia is characterized by gelatinous basidiomata that are resupinate to substipitate with a hairy upper surface, cylindrical to clavate basidia with three transverse septa and oil guttules, and hyaline, thin-walled, allantoid basidiospores [1,2,5]. Members of this genus are saprotrophic, commonly growing on dead trees, stumps, fallen trunks and branches, and other decaying wood, and are broadly distributed across tropical, subtropical, and temperate regions worldwide [2,6]. Several Auricularia species are highly valued as edible mushrooms and contribute significantly to the economy, particularly in China and other East Asian countries, e.g., A. cornea and A. heimuer [7,8]. Auricularia species are cultivated as an important food source due to their high protein, carbohydrate, and mineral contents, as well as their low fat content [9–11]. Medicinally, they possess anticancer, anticoagulant, antimicrobial, cholesterol-lowering, detoxifying, and hypoglycemic properties, providing various health benefits to humans [12–14]. Traditional identification of Auricularia species is mainly based on morphological characteristics [3,15–17]. Based on morphological characteristics, Auricularia has been classified into five species complexes: A. auricula-judae, A. cornea, A. delicata, A. fuscosuccinea, and A. mesenterica complexes [2,16]. However, identification is challenging due to the high phenotypic variability across broad geographic ranges, the influence of varying environmental conditions, and the difficulty in distinguishing morphological traits at different developmental stages among closely related Auricularia species. Thus, it is essential to identify Auricularia species by applying DNA techniques and phylogenetic analyses. Phylogenetic analyses combining ITS, nrLSU, and rpb2, or using ITS alone, have recently been employed for the identification of Auricularia [2,3,18,19]. These analyses have shown that both the A. auricula-judae and A. mesenterica complexes form monophyletic groups, whereas the other species complexes are polyphyletic [2]. Currently, 37 species of Auricularia have been accepted [2]. DNA sequences were successfully obtained from 31 Auricularia species, whereas sequencing failed for six other species (A. eburnea, A. eminii, A. hainanensis, A. minor, A. papyracea, and A. xishaensis).
     Thailand is considered a biodiversity hotspot, particularly in its northern regions. According to previous studies, northern Thailand has been the source of the majority of newly reported mushroom species [20–24]. Auricularia has been called by various common names, such as ear mushrooms, wood ear, Jew's ear, jelly fungus, and “Hed-hoo-noo” in Thai. The first scientific investigations of Auricularia were carried out by Johannes Schmidt during his 1899–1900 expedition to Siam [24]. Prior to this study, 13 Auricularia species, namely A. asiatica, A. auricula-judae (syn. A. auricula and A. auriculalis), A. cornea, A. delicata, A. fibrillifera, A. fuscosuccinea, A. mesenterica, A. ornata, A. peltate, A. nigricans (syn. A. polytricha), A. tenuis, A. thailandica, and A. villosula have been reported in Thailand [3,18,19,25–28]. In this study, we found three Auricularia specimens during our investigations of mushrooms in northern Thailand from 2024 to 2025. This study aimed to identify these specimens using both morphological characteristics and molecular data. Based on these analyses, the specimens were identified as A. sinodelicata, a species not previously reported in Thailand. A phylogenetic tree based on combined ITS, nrLSU, and rpb2 sequence data was constructed to confirm the phylogenetic positions of this species. Furthermore, the taxonomic status of Auricularia in Thailand is discussed. An updated key and checklist of Auricularia species recorded in the country are also provided.

2. MATERIALS AND METHODS

2.1 Sample Collection
     Auricularia specimens were collected from community forests in Chiang Mai Province, Thailand, during the rainy seasons of 2024 and 2025. Photographs of the fresh specimens were taken in the field using a Canon EOS 700D digital camera (Canon Inc., Japan). Basidiomata were collected and placed in plastic boxes for transport to the Center of Excellence in Microbial Diversity and Sustainable Utilization at Chiang Mai University for laboratory analysis. After that, fresh specimens were dried in a hot air oven at 45°C until completely dry [29]. Dried specimens were then kept in a plastic zip-lock bag and deposited at the Chiang Mai University Biology Department’s Herbarium (CMUB) and Herbarium of Sustainable Development of Biological Resources (SDBR), Faculty of Science, Chiang Mai University, Chiang Mai, Thailand.

2.2 Morphological Observation
     Macromorphological data were recorded from fresh specimens within 24 hours of collection. Color codes followed Kornerup & Wanscher [30]. Micromorphological data were obtained from dried specimens mounted in 5% KOH, Congo red solution, or Melzer’s reagent, and characteristics were examined under a light microscope (Nikon Eclipse Ni U, Tokyo, Japan). Size data of the micromorphological structures (e.g., basidiospores, and basidia) are based on at least 25 measurements of each structure using the Tarosoft (R) Image Frame Work program. Basidiospore statistics are expressed as (a–)b–c(–d), where ‘a’ and ‘d’ are the extreme values, and ‘b–c’ is the range comprising 95% of all values. Q represents the ratio of length to width of each basidiospore. The terminology of the microscopic features followed Lowy [1] and Wu et al. [2].

2.3 DNA Extraction and PCR Amplification
     DNA was extracted directly from fresh specimens using a DNA Extraction Mini Kit (FAVORGEN, Taiwan) following the manufacturer’s protocol. The ITS, nrLSU, and rpb2 loci were amplified by polymerase chain reaction (PCR) with ITS5/ITS4 [31], LR0R/LR5 [32], and b6F/b7.1R [33] primer pairs, respectively. The amplification of these three loci was performed in separate PCR reactions under the same conditions, consisting of an initial denaturation at 95°C for 5 min, followed by 35 cycles of denaturation at 95°C for 30 s, annealing at 52°C for 1 min (ITS and nrLSU) or at 55°C for 1 min (rpb2), extension at 72°C for 1 min, and a final extension at 72°C for 10 min, using a peqSTAR thermal cycler (PEQLAB Ltd., UK). PCR products were examined on 1% agarose gels stained with ethidium bromide under UV light. The amplified products were then purified using the NucleoSpin® Gel and PCR Clean-up Kit (Macherey-Nagel, Germany) according to the manufacturer’s instructions. The purified PCR products were directly sequenced using a genetic analyzer at 1^st Base Company (Kembangan, Malaysia) with the same PCR primers.

2.4 Sequence Alignment and Phylogenetic Analyses
     The generated sequences were assembled and manually edited using Sequencer version 5.4.6 (Gene Codes Corp., USA) to obtain consensus sequences, and sequence quality was assessed by inspecting chromatograms and trimming low-quality regions at both ends. The consensus sequences were analyzed using BLAST searches against the GenBank database (http://blast.ncbi.nlm.nih.gov, accessed on 30 October 2025). Sequences from this study, along with relevant sequences from previous studies and the GenBank database, were selected for analysis as listed in Table 1. Multiple sequence alignment was performed with MUSCLE [34] using default settings and improved where necessary using BioEdit version 6.0.7 [35].
     Phylogenetic analysis was performed using a combined dataset of ITS, nrLSU, and rpb2 sequences. A phylogenetic tree was constructed using maximum likelihood (ML) and Bayesian inference (BI). Exidia subsaccharina BJFC 036778 and Heteroradulum australiense 20180515-26 were set as outgroup. The ML analysis was performed using 25 categories and 1000 bootstrap (BS) replicates with the GTRCAT model of nucleotide substitution on RAxML-HPC2 version 8.2.12 [36] at the CIPRES web portal. The GTR+I+G model was selected as the best model. BI analysis was performed using the MrBayes version 3.2.6 software [37]. The best substitution models for each gene were determined using the Akaike Information Criterion (AIC) in jModelTest 2.1.10 [38]. For BI analysis, six simultaneous Markov chains were run for one million generations with random initial trees, with sampling every 1,000 generations. The first 20% of generated trees, representing the burn-in phase of the analysis, were discarded, while the remaining trees were used to calculate PP in the majority-rule consensus tree. The Bayesian posterior probabilities (PPs) were subsequently calculated. The phylogenetic trees were visualized using FigTree version 1.4.0 [39].

3. RESULTS

3.1 Sample Collection and Morphological Study
     Three Auricularia specimens were collected from a deciduous community forest in Chiang Mai Province, Thailand, during the 2024 and 2025 rainy seasons. These specimens were deposited in the herbarium under the accession numbers CMUB40117, SDBR-CMUNK1948, and SDBR-CMUNK2435. Based on morphological characteristics, these specimens were classified in the A. delicata complex and showed similarity to A. sinodelicata [2]. Therefore, molecular phylogenetic analysis was used for species identification.

3.2 Phylogenetic Analyses
     The ITS, nrLSU, and rbp2 sequences of each specimen were deposited in the GenBank database (Table 1). The ITS and nrLSU sequences obtained in this study showed 99.63% and 99.56% similarity to the ITS and nrLSU of A. sinodelicata Dai 13926 (holotype specimen), respectively. For phylogenetic analyses, the combined ITS, nrLSU, and rbp2 sequence dataset comprised 124 taxa, and the aligned dataset consisted of 2,513 characters, including gaps (ITS: 1–620; nrLSU: 621–1,979; and rbp2: 1,980–2,513). The best-scoring RAxML tree was established with a final ML optimization likelihood value of -13622.3386. Accordingly, the matrix contained 680 distinct alignment patterns, with 33.99% of characters undetermined or missing. The estimated base frequencies were recorded as follows: A = 0.2462, C = 0.2373, G = 0.2801, T = 0.2362; substitution rates AC = 1.5184, AG = 3.5943, AT = 1.7454, CG = 1.4857, CT = 7.5876, GT = 1.0000. The tree length and gamma distribution shape parameter alpha values were equal to 1.6454 and 0.1339, respectively. In addition, the BI analysis converged with a final average standard deviation of split frequencies of 0.008772 after one million MCMC generations. The topologies of the phylogenetic trees obtained from ML and BI analyses were similar. Therefore, a phylogenetic tree generated from the ML analysis is shown in Figure 1. The phylogenetic tree obtained in this study was consistent with the findings of Wu et al. [2]. In addition, the phylogenetic clades were separated similar to Wu et al. [2]. The result indicated that both the A. auricula-judae and A. mesenterica complexes presented as monophyletic lineages, while other Auricularia species complexes showed polyphyletic lineages. Three Auricularia specimens obtained in this study clustered within the A. sinodelicata clade. Therefore, they were identified as A. sinodelicata. Auricularia sinodelicata was closely related to A. delicata and A. lateralis.
     Previously published sequences of six Auricularia species (A. asiatica, A. cornea, A. delicata, A. fibrillifera, A. thailandica, and A. villosula) collected in Thailand were also included in the phylogenetic analyses. The results showed that all species formed well-supported clades corresponding to their respective taxa, except for A. delicata. Based on phylogenetic evidence, Thai specimens previously identified as A. delicata [18] were clustered within the A. sinodelicata clade. Moreover, their ITS sequences showed 96.39%–97.72% similarity to A. delicata P 14 (epitype specimen) and 98.30%–99.43% similarity to A. sinodelicata Dai 13926 (holotype specimen), respectively. Therefore, the Thai specimens previously identified as A. delicata should be reclassified as A. sinodelicata based on phylogenetic evidence.

3.3 Taxonomic Description
Auricularia sinodelicata Y.C. Dai & F. Wu, Journal of Fungi 7(11): e933 (2021) Figure 2.
MycoBank number: MB 825100

     Basidiomata: Gelatinous when fresh, fawn to reddish brown (8E6) or greyish orange (6B5) to light brown (7C5), solitary or caespitose, sessile or substipitate; pileus discoid or auriculate, sometimes with lobed margin, projecting up to 7 cm, 1–2.5 mm thick, with a hymenial layer 0.15–0.25 mm thick, reddish brown (8E5) to dark brown (8F4) when dry; upper surface scantly pilose; hymenophore surface conspicuously porose-reticulate.
     Internal features: Medulla indistinctly present near the hymenium; crystals absent; abhymenial hairs with a slightly swollen base, hyaline, thick-walled, with a wide or narrow septate lumen, apical tips acute or obtuse, single, 25–80 × 5–8 µm; hyphae with clamp connections, 1–5 µm wide. Basidia clavate, transversely 3-septate, with oil guttules, 30–40 × 4–5 µm, sterigmata rarely observed. Cystidioles absent. Basidiospores (9.8–)10–12(–12.2) × (4–)4.4–5.0(–5.5) µm (n = 50), Q = 2.2–2.4, allantoid, hyaline, smooth, thin-walled, usually with one to three large guttules.
     Material examined: THAILAND, Chiang Mai Province, Mae Taeng District, Pa Pae, 19°06'48"N 98°42'45"E, elevation 854 m, on decaying wood in a deciduous community forest, 23 May 2024, J. Kumla & N. Suwannarach, SDBR-CMUNK1948; Mae Wang District, 18°37'30"N 98°48'59"E, elevation 330 m, on decaying wood in a deciduous community forest, 10 August 2025, J. Kumla & N. Suwannarach, CMUB40117 = SDBR-CMUNK2349; Mueang Chiang Mai District, 18°43'49"N 98°54'50"E, elevation 341 m, on decaying wood in a deciduous community forest, 1 October 2025, N. Suwannarach, SDBR-CMUNK2435.
     Habitat and known distribution: Fruiting on dead trees, fallen trunks and branches, and decaying wood. Known from China [2], India [40], and Thailand [This study].

4. DISCUSSION

     Auricularia species are currently identified based on both morphological characteristics and molecular evidence. In this study, A. sinodelicata was discovered in northern Thailand and identified using an integrated approach involving morphological examination and multi-gene phylogenetic analyses. Prior to this study, A. sinodelicata was found in China [2] and India [40]. Therefore, this is the first report of this species in Thailand and Southeast Asia. Morphologically, A. sinodelicata is similar to A. delicata and A. lateralis in having a porose-reticulate hymenophore, as shown by phylogenetically closely related taxa, but they are different lineages. Auricularia sinodelicata has shorter basidia (30–45 × 4–5.5 µm) than A. delicata (48–65 × 4–6 µm) [1,2,16, This study]. Auricularia lateralis has larger basidiospores (12.9–14.2 × 5.2–6 µm) than A. sinodelicata (10–12 × 4.3–5.1 µm) [2]. Based on their distribution, A. delicata and A. lateralis have been reported only from West Africa [1,16] and China [2], respectively.
     The first scientific investigations of Auricularia were carried out by Johannes Schmidt during his 1899–1900 expedition to Siam [24]. Subsequently, field reports documented the occurrence of A. delicata, A. fuscosuccinea, A. mesenterica, and A. polytricha (currently known as A. nigricans) in Thailand, although molecular data were lacking and herbarium material was insufficient [25–28]. In 2011, the “Checklist of Mushrooms (Basidiomycetes) in Thailand” listed Thai mushrooms belonging to the phylum Basidiomycota, including eight Auricularia species, namely A. auricula-judae, A. delicata, A. fuscosuccinea, A. mesenterica, A. ornata, A. peltate, A. nigricans, and A. tenuis [25]. However, these species were listed based on morphological characteristics, while molecular data and herbarium material are still lacking. In 2015, the first investigation of Auricularia in Thailand was conducted using both morphological and molecular phylogenetic analyses, and A. thailandica was proposed [3]. In 2017, four Auricularia species, including A. asiatica, A. cornea, A. delicata, and A. villosula, were reported in Thailand [18,41–43]. However, specimens previously identified as A. delicata were placed within the A. sinodelicata clade based on multi-gene phylogenetic analyses (Figure 1), indicating that Thai specimens previously identified as A. delicata should be reclassified as A. sinodelicata based on phylogenetic results. The morphological characteristics of these species should be re-examined to confirm that they are A. sinodelicata. Subsequently, A. fibrillifera was reported in Thailand in 2024 [19]. In this study, A. sinodelicata was collected and identified. Therefore, a total of 14 Auricularia species have been listed in Thailand. However, molecular data and Thai specimens for the remaining eight species (A. auricula-judae, A. delicata, A. fuscosuccinea, A. mesenterica, A. ornata, A. peltate, A. nigricans, and A. tenuis) are still lacking and require further investigation and taxonomic confirmation. Consequently, an updated list of Auricularia species recorded in Thailand, based on both morphological and molecular evidence, is presented in Table 2. Additionally, a key to accepted species of Auricularia in Thailand, modified from Wu et al. [2], is provided.

5. CONCLUSIONS

     In this study, A. sinodelicata collected from northern Thailand were identified based on a combined assessment of their morphological characteristics and multi-gene phylogenetic analysis. This species is a new record for Thailand and a new distribution record in Southeast Asia. The findings of this study indicate that 14 Auricularia species have been recorded in Thailand, of which both morphological and molecular data support six species, and the remaining eight require new specimen collections and molecular evidence for taxonomic confirmation. Moreover, an updated key and checklist of Auricularia species in Thailand are provided. This study could serve as an inspiration, motivating researchers to investigate the distribution and ecology of Auricularia in Thailand, Asia, and worldwide. Further research is required to investigate the cultivation potential, nutritional composition, and medicinal properties of A. sinodelicata. Furthermore, climate change should be considered in future research on Auricularia in Thailand, given its potential effects on distribution, habitat, and species diversity.

ACKNOWLEDGEMENTS

     This research was financially supported by Chiang Mai University, Thailand.

AUTHOR CONTRIBUTIONS

Jaturong Kumla: Conceptualization, Data curation, Investigation, Methodology, Software, Validation, Writing - Original draft preparation. Nakarin Suwannarach: Conceptualization, Formal analysis, Funding acquisition, Methodology, Project administration, Resources, Supervision, Validation, Visualization, Writing- Reviewing and Editing.

CONFLICT OF INTEREST STATEMENT

     The authors declare that they hold no competing interests.

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José R. Fuentes-Castillo, Flor N. Rivera-Orduña, Juan C. Cancino-Díaz, Juan A. Cruz-Maya and Janet Jan-Roblero

Vol.52 No.5 (September 2025)
Research Article View: 566 Download: 204

Complete Mitochondrial Genome of a Karst-dwelling Gecko, Gekko liboensis (Zhou and Li, 1982)
DOI: 10.12982/CMJS.2025.048.

Jing Cao, Phuping Sucharitakul, Fang Yan, Chatmongkon Suwannapoom and Siriwadee Chomdej

Vol.52 No.4 (July 2025)
Research Article View: 614 Download: 181

Addition to Paraleptosphaeria species (Leptosphaeriaceae, Pleosporales): Paraleptosphaeria senecionis sp. nov., associated with Asteraceae
DOI: 10.12982/CMJS.2025.014.

Ausana Mapook, Erio Camporesi and Kevin David Hyde

Vol.52 No.2 (March 2025)
Research Article View: 813 Download: 216